Four New Species of Paspalum (Poaceae, Paniceae) from Central Brazil, and Resurrection of an Old One
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Systematic Botany (2008), 33(2): pp. 267–276 © Copyright 2008 by the American Society of Plant Taxonomists Four New Species of Paspalum (Poaceae, Paniceae) from Central Brazil, and Resurrection of an Old One Gabriel H. Rua,1,4 José F. M. Valls,1 Dalva Graciano-Ribeiro,2 and Regina C. Oliveira3 1EMBRAPA Recursos Genéticos e Biotecnologia, Parque Estação Biológica, Final W-5 Norte, Caixa Postal 02372, CEP 70770-900 Brasília, DF, Brazil 2Universidade de Brasília, Instituto de Ciências Biológicas, Departamento de Botânica, Campus Universitário, Asa Norte, Caixa-Postal 04457, 70910-900 Brasília, DF, Brazil 3Herbário Dárdano de Andrade Lima, Universidade Federal Rural do Semi-Árido, Caixa Postal 137, CEP 59600-970, Mossoró, RN, Brazil 4Author for correspondence, present address: Cátedra de Botánica Agrícola, Facultad de Agronomía, Universidad de Buenos Aires, Av. San Martín 4453, C1417DSE Buenos Aires, Argentina ([email protected]) Communicating Editor: Gregory M. Plunkett Abstract—Four new Brazilian species from the genus Paspalum are described and illustrated: P. phaeotrichum, P. vexillarium, P. veredense, and P. clipeum. Paspalum phaeotrichum is an annual with no obvious affinity to any known species of Paspalum, although it shares several characters with species of both P. subg. Ceresia and the ’Bertoniana’ group. Paspalum vexillarium is presumably related to P. ceresia, with which it has been confused. Paspalum veredense shows affinities with both P. ellipticum and P. erianthoides. Paspalum clipeum is probably related to annual species of the ’Plicatula’ group, although it lacks the dark brown upper florets typical of that group. Moreover, P. spissum, a species currently considered as a synonym under P. carinatum, is recognized on the basis of vegetative morphology, phenology and habitat. Keys to the treated species are provided when relevant, as well as a distribution map. Resumo—Quatro novas espécies de Paspalum do Brasil são descritas e ilustradas: P. phaeotrichum, P. vexillarium, P. veredense,eP. clipeum. Paspalum phaeotrichum é uma espécie anual sem afinidades obvias com outras espécies do gênero, embora compartilhe vários caracteres com P. subg. Ceresia e com o grupo ’Bertoniana’. Paspalum vexillarium é supostamente relacionada a P. ceresia, com a qual tem sido confundida. Paspalum veredense tem afinidades com P. ellipticum e P. erianthoides. Paspalum clipeum é provavelmente relacionada às espécies anuais do grupo ’Plicatula’, embora não apresente os antécios superiores de cor marrom escura, típicos desse grupo. Além disso, P. spissum, uma espécie atualmente considerada como sinônimo de P. carinatum, é reabilitada com base na morfologia vegetativa, fenologia e habitat. São fornecidas chaves para as espécies tratadas, quando relevante, assim como um mapa de distribuição. Keywords—Brazilian flora, Cerrado, grasses, Panicoideae, taxonomy. Paspalum L. comprises about 350 species of grasses mostly Chase was examined at BAA and SI for comparison. Leaf samples for distributed in tropical and warm-temperate areas of the anatomical sections from herbarium specimens were rehydrated accord- ing to the method described by Voster (1990) with few modifications. Americas (Clayton and Renvoize 1986; Zuloaga and Morrone Cross sections were obtained using a Ranvier microtome, stained with 2005; Denham 2005). They are recognized by their dorsiven- Alcian blue and safranin 1% and mounted using a synthetic resin (Paiva tral raceme-like partial inflorescences, by their plano-convex et al. 2006). Anatomical micrographs were obtained using a Zeiss Ax- spikelets with the upper lemma adaxial, and, in most cases, ioscop MC80DX microscope. Micrographs of the upper lemma surface were taken using a Zeiss DSM 962 scanning electron microscope (SEM). by the lacking of the lower glume. Within the Panicoideae, the genus Paspalum belongs to a clade characterized by a basic chromosome number of x = 10 (Giussani et al. 2001) and TAXONOMIC TREATMENT is related to other genera having an NADP-ME photosyn- thetic pathway. Paspalum phaeotrichum Valls, G. H. Rua, Graciano-Ribeiro, Species of Paspalum are, in large part, responsible for much & R. C. Oliveira, sp. nov.—TYPE (here designated): of the biodiversity of grassland ecosystems in South America, BRAZIL. Bahia: Mun. Riachão das Neves, entrada de which are strongly threatened by the expansion of agricul- fazenda ca. 20 km N de Barreiras pela estrada antiga, 2 ture. Among them, the Brazilian Cerrado is certainly a km N da ponte sobre o Rio Branco, 31 Mar 1983, J. F. M. hotspot of grass diversity (Warming 1973; Mendonça et al. Valls, A. Krapovickas, R. F. A. Veiga, & G. Pereira-Silva 7049 1998). A number of new grass genera and species have been (holotype: CEN!; isotypes: BAA!, L!, RB!, SI!, US!). described from this region in recent years, including several species of Paspalum (Burman 1980; Davidse and Filgueiras A omnibus speciebus notis distincta, tamen P. lilloi Hack. 1993; Filgueiras and Davidse 1994; Filgueiras 1995; Davidse flosculo molli, laminis foliorum costatis aliquantum similis, et al. 2001; Filgueiras et al. 2001, Morrone and Zuloaga 2003). sed habitu annuo, omnibus partibus multo minoribus differt. While identifying grasses collected in Cerrado areas of cen- Annuals. Culms 18–28 cm tall, 0.8–0.9 mm diam, erect to tral Brazil, four new species of Paspalum were found. They decumbent, freely branching, some culms proximally creep- are here described, illustrated, and compared with related ing, with branches rooting at the base, the main axis coming species. Moreover, P. spissum, a species currently considered to flower; internodes glabrous, the basalmost purple, other- a synonym of P. carinatum, is recognized on the basis of veg- wise pale; nodes glabrous, purple-tinged. Leaf sheaths etative morphology, phenology and habitat. keeled, striate, long-ciliate at the top of margins, otherwise glabrous, the proximal sheaths 1–1.5 cm long, purple-tinged, the distal sheaths up to 6 cm long. Ligules 0.5–0.7 mm long, MATERIALS AND METHODS membranous, acute, glabrous, erose, without a pseudoligule. Descriptions and illustrations were based on field collections and her- Blades 4.5–5.5 cm long, ca. 2 mm wide, lanceolate-acuminate, barium material (CEN, IBGE, UB). Andean material of P. ceresia (Kuntze) flat, the base narrower than the leaf sheath, sharply ribbed 267 268 SYSTEMATIC BOTANY [Volume 33 FIG. 2. Leaf blade anatomy in cross section, general view (left) and detail of a portion (right). A, B. Paspalum phaeotrichum (Valls et al. 7049). C, D. Paspalum vexillarium (Amaral et al. 28). E, F. Paspalum veredense (Rua et al. 695). FIG.1. Paspalum phaeotrichum (from Valls et al. 7049). A. Habit. B. Spikelet, adaxial view. C. Spikelet, abaxial view. D. Upper glume, adaxial overlapping marginal flaps; lodicules 2; stamens 3, anthers view. E. Lower lemma, abaxial view. F. Upper floret, adaxial view. G. 2.1–2.2 mm long, yellowish; stigmas 2, pale, plumose. Cary- Upper floret, abaxial view. H. Caryopsis, hilum side. I. Caryopsis, em- bryo side. J. Caryopsis, lateral side. K. Portion of the rachis, abaxial view. opsis 1.8 mm long, ca. 1 mm wide, plano-convex, obovate, slightly emarginate at apex, pale; hilum ca. 0.8 mm long., adaxially; the margins with a few proximal cilia, otherwise linear-elliptical. Figure 1. glabrous; cauline blades reduced, the most distal ca. 1 cm Leaf Anatomy—(Fig. 2A, B). Leaf blade expanded in cross long. Peduncles 4.5–9 cm long, compressed, glabrous, pale to section, sharply ribbed/furrowed adaxially. Midrib incon- purple-tinged. Inflorescences exserted; main axis not pro- spicuous, with a single first-order vascular bundle. Cuticle longed above the racemes; ‘racemes’ 2–4 cm long, 2, conju- thin. Adaxial epidermis with typical epidermal cells, bicellu- gate, ascending to divaricate; pulvini hirsute; rachis of the lar microhairs, 3–5 fan-shaped bulliform cells, stomata, and racemes ca. 0.4 mm wide, glabrous, flexuous, triquetrous, narrowly 3-winged, ending in a terminal spikelet, the nerves green, otherwise pale; spikelets 4.0–4.1 mm long, 1.1–1.4 mm wide, narrowly ovate, solitary, sessile, loosely 2-seriate, de- ciduous at maturity, dorsiventrally compressed. Lower glumes lacking. Upper glumes as long as the spikelets, ca. 2.1 mm wide, ovate, membranous-hyaline, pale, 7-nerved, the two outer nerve pairs close to the margins, the proximal half convex, corrugate, densely pilose with pale hairs inter- mingled with longer and stiffer, golden-brown cilia up to 3 mm long; the distal half depressed, somewhat ornamented with longitudinal purple stripes, pilose along the lateral nerves, otherwise glabrous, the involute margins covering the margins of the lower lemma; the base widely attenuate, the apex acute but somewhat rounded at the very tip. Lower lemmas ca. 4 mm long, 1 mm wide, narrowly ovate, flat, acute at apex, 5-nerved, the lateral nerves close to the mar- gins, densely pilose at the proximal portion. Upper florets 3.1–3.3 mm long, 1–1.1 mm wide, narrowly ellipsoidal, flat- tened, hyaline, pale, glabrous, the upper lemma faintly 5-nerved, slightly shorter that the palea and not enclosing the FIG. 3. Distribution map of Paspalum phaeotrichum, P. vexillarium, P. palea apex, so that the floret remains open, the palea with veredense, P. clipeum, and P. spissum. 2008] RUA ET AL.: NEW PASPALUM FROM BRAZIL 269 silicified cells. Mesophyll consisting of radial chlorenchyma finity with any known species of Paspalum. Morphologically cells. Three orders of vascular bundles present, each sur- it seems allied to species of P. subg. Ceresia (Pers.) Rchb., rounded by a single bundle sheath of relatively large, thin- because of its soft, distally gaping upper floret, and the hair walled parenchyma cells, the bundles separated by 1 or 2 pattern in the basal portion and margins of the upper glume. parenchyma cells. Sclerenchyma both adaxial and abaxial However, the rachis of the ‘racemes’ are slender, triquetrous, girders along ribs, but only abaxially at intercostal regions. narrowly 3-winged, with the lateral margins not distinctly Distribution and Habitat—(Fig.